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Soft Corals (Octocorallia: Alcyonacea) from Southern Taiwan. II. Species

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Soft Corals (Octocorallia: Alcyonacea) from Southern Taiwan. II. Species Zoological Studies 43(3): 548-560 (2004) Soft Corals (Octocorallia: Alcyonacea) from Southern Taiwan. II. Species Diversity and Distributional Patterns Yehuda Benayahu1,*, Ming-Shiou Jeng2, Shimrit Perkol-Finkel1 and Chang-Feng Dai3 1Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Ramat Aviv, Tel Aviv 69978, Israel 2Institute of Zoology, Academia Sinica, Taipei, Taiwan 115, R.O.C. 3Institute of Oceanography, National Taiwan University, Taipei, P.O. Box 23-13, Taiwan 106, R.O.C. (Accepted June 28, 2004) Yehuda Benayahu, Ming-Shiou Jeng, Shimrit Perkol-Finkel and Chang-Feng Dai (2004) Soft corals (Octocorallia: Alcyonacea) from southern Taiwan. II. species diversity and distributional patterns. Zoological Studies 43(3): 548-560. The geographic setting of Taiwan, between the West Pacific Ocean and East China Sea and at the crossroads of the Philippine-Japan I. Arc, has produced reefs of special biogeographical inter- est. It has been suggested that Taiwan may have served as a“stepping-stone”for the dispersal of shallow reef organisms. Soft corals from Nanwan Bay and Green I. (Lutao in Chinese), southern Taiwan, were studied dur- ing 1994 and 1998. Scuba collections were carried out to a depth of 33 m. Approximately 230 samples were collected, encompassing a variety of species found on these reefs. Visual estimates were made of the under- water abundance of species. The collection yielded 69 species of the families Helioporidae, Clavulariidae, Tubiporidae, Alcyoniidae, Nephtheidae, Xeniidae, and Briareidae. These included 1 new species and 43 new zoogeographical records for Taiwan. Among the 22 listed genera, the survey recorded 7 for the 1st time in Taiwanese reefs. The findings confirm the high soft-coral diversity of these reefs. The shallow reefs of Nanwan Bay are densely inhabited by species of the family Alcyoniidae. In contrast, the deep reefs are characterized only by sporadic colonies of Alcyoniidae, but abundant assemblages of azooxanthellate members of the fami- lies Nephtheidae and Nidaliidae. Soft corals of the family Xeniidae are abundant on the reefs of Green I. at a depth range of 3~10 m. Interestingly, species of the family Xeniidae were rarely observed at Nanwan Bay. The coral reefs of Taiwan and Japan are closely linked by the northward-flowing Kuroshio Current, which brings water and larvae from the reefs of the South China Sea. Therefore, we compared the generic affiliation and abundance estimates of the Alcyoniidae between southern Taiwan and the Ryukyu Archipelago (Japan), and found a close resemblance between these 2 reef areas. http://www.sinica.edu.tw/zool/zoolstud/43.3/548.pdf Key words: Octocorallia, Taiwan, Coral reefs, Species diversity, Biogeography. Taiwan, a continental island with several off- crossroads of the Philippine-Japan I. Arc, has pro- shore islets, has coral reefs that sit at the northern duced reefs of special biogeographical interest. edge of the Southeast Asian center of biodiversity This has led to the suggestion that Taiwan may (Fujiwara et al. 2000). These reefs are known for have served as a“stepping-stone”for the north- their flourishing fringing reef communities (Wells ward and eastward dispersal of shallow reef 1988). The Taiwan Strait, situated to the west organisms (Dai 1991a). between Taiwan and mainland China, is a shallow The octocoral fauna of Taiwan has been the channel, as opposed to the eastern region where subject of several studies. An early collection of the submarine topology drops steeply close to the soft corals conducted by Utinomi in 1938, resulted nearby Ryukyu Archipelago (Chen 1999). The in the description of 3 new species: Clavularia geographic setting of Taiwan, between the West racemosa, Anthelia formosana, and Asterospi- Pacific Ocean and East China Sea and at the cularia laurae (Utinomi 1950a b 1951). This col- *To whom correspondence and reprint requests should be addressed. Tel: 972-3-6409090. Fax: 972-3-6409403. E-mail: [email protected] 548 Benayahu et al. -- Living Soft Corals of Southern Taiwan 549 lection was carried out mainly along the southern MATERIALS AND METHODS coast of Taiwan, and yielded 21 octocoral species of the families Clavulariidae, Tubiporidae, Collecting trips were conducted in 2 regions Xeniidae, Asterospiculariidae, Alcyoniidae, (Fig. 1): Kenting National Park, Nanwan Bay in Nephtheidae, Helioporidae, and Briareidae southern Taiwan (21 55'N; 120 45'E) during (Utinomi 1959). Later, Verseveldt (1982a) June-July 1994, Nov. °1998, and Mar.° 2004; and described an additional new species, Siphono- the offshore Green I. (Lutao in Chinese), southeast gorgia lobata (family Nidaliidae), from Taiwan. The of Taiwan (22 40'N; 121 29'E) during June-July 18 species of gorgonians listed for southern 1994. In Nanwan° Bay, collections° were made on Taiwan, all of which were found to be new records the fringing reefs including Banana Bay and for the region (Chen and Chang 1991), are like- Wanitun, mostly to a maximal depth of 12 m, and wise notable, but are beyond the scope of the cur- on the submerged offshore Sea Mount reef, down rent paper. Fujiwara et al. (2000) reported that to 31 m. At Green I., collections were made along species diversity of the Taiwanese reefs is relative- the western and southwestern sides down to a ly high, with 70 soft-coral and gorgonian species. depth of 31 m. The reefs were mostly reached by Dai (1991a b) demonstrated that soft corals in boat, and careful examination of a variety of niches Taiwan are abundant in the south, mainly on the in the different sites was carried out by scuba div- western side of Nanwan Bay. The reefs of Kenting ing. Approximately 230 samples were collected, National Park in southern Taiwan are very rich, encompassing the variety of species found on the with extensively developed soft-coral communities. reefs. Underwater abundance estimates of the They dominate the benthos, comprising in some genera were made visually and divided into 4 cate- sites up to 70% of the total coral cover (Dai 1993). gories: rare, sporadic, abundant, and dominant In these reefs, species of the genera Lobophytum, (see also Benayahu 1995). Prior to collection, Sarcophyton, and Sinularia are highly abundant most of the colonies were in situ photographed, and found even in habitats exposed both to strong using a Nikonos V camera with a close-up attach- tidal currents and typhoons (Dai 1988 1991b). A ment. Samples were fixed in 4% formalin in sea- checklist of the soft corals from southern Taiwan (Dai 1991a) included 42 species and an additional 7 species that remained unassigned. Among the 26°N N identified species of this list, Utinomi (1959) had already reported 13 species, and therefore 29 species appeared to be new records for the region. At present, there is a gap in the knowledge of soft corals from other regions of Taiwan. 25°N Intrigued by the early taxonomic reports on the soft corals of Taiwan (Utinomi 1950a b 1951 1959), their high abundance, and ecological impor- tance (Dai 1988 1990 1991a b 1993; Fujiwara et 24°N al. 2000), we conducted a comprehensive soft- Penghu Is. TAIWAN coral survey there. The present paper reports on reef sites located in both the East China Sea and the Pacific side of southern Taiwan. It provides a 23 N systematic list of soft-coral species, as well as ° Green I. information on their abundance and distributional Green I. patterns on the reefs. The blue coral Heliopora coerulea is abundant in some of the surveyed Lanyu sites, and therefore it is also included in the study. 22°N Since the coral reefs of Taiwan and Japan are closely linked, we also compared the studied soft- coral fauna to those of reefs of the adjacent south- ern Ryukyu Archipelago and assessed their bio- Nanwan geographic affinities. 21 N ° 120°E 121°E Fig. 1. Map of Taiwan with study sites indicated by asterisks. 550 Zoological Studies 43(3): 548-560 (2004) water, rinsed in fresh water after 24 h, and then Zoology, Tel Aviv University, Israel. Some mem- transferred to 70% ethyl alcohol. Sclerites were bers of the families Alcyoniidea, Nephtheidae, obtained by dissolving the tissues in 10% sodium Nidaliidae, and Xeniidae are still being examined. hypochlorite. Identification of the Alcyoniidae species of the collection was in great part facilitated by compar- RESULTS isons with permanent sclerite preparations from type material kept in the Zoological Museum, The examined material yielded 69 species, of Department of Zoology, Tel Aviv University, Israel which 1 is a new species (Benayahu and Perkol- (ZMTAU) and at the Nationaal Natuurhistorisch Finkel 2004) and 43 are new zoogeographical Museum, formerly Rijksmuseum van Natuurlijke records for Taiwan (Table 1). Underwater pho- Historie, Leiden, The Netherlands. All material is tographs of some of the soft corals collected in the housed in the Zoological Museum, Department of current study are presented as indicated below. Table 1. List of the species of Octocorallia of the orders Helioporacea Bock, 1938 and Alcyonacea Lamouroux, 1816 from the reefs of Taiwan with indication of Museum inventory numbers (ZMTAU Co) and previous references. NR, new record; 1, Utinomi 1959; 2, Dai 1991a; * occurrence of members of the family Alcyoniidae in the southern Ryukyu Archipelago, Japan following Benayahu 1995 2002; ** see Benayahu and Perkol-Finkel 2004 Classification References Classification References Helioporidae Blainville, 1830 30252 Genus Heliopora Moseley, 1786 Lobophytum sarcophytoides Moser, 1919 2 Heliopora coerulea
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